No. 105 MARCH 1970
RESOURCE MANAGEMENT REPORT
DEPARTMENT OF LANDS AND FORESTS
HON. RENE BRUNELLE G. H. U. BAYLY
Minister Deputy Minister
RESOU RCE MANAG EMENT REPORT
NO. 105 ~" MftitC H, 1970
PAGE M f . INSTRUCTIONS
1 In heading, 6th line, delete "Lake Simcoe District".
1 No. 1, (b) , 5th line, after "white" insert "pine, basswood, white
2 No. 1, (d), 1st paragraph, 3rd line, after "rather" insert "than".
2 No. 1, (e), 2nd paragraph, 1st line, delete "the", insert "its".
4 No. 3, (a), 4th line, delete "growth", insert "grown".
5 No. 3, (d) , last line, after "moist" insert "forest".
6 No. 3, (d) , 2nd paragraph, 13th line, delete "shade", insert
10 No. 5, (a), 4th paragraph, last line, delete "germinating",
13 No. 6, (b) , 1st line on page, not a new paragraph.
14 No. 7, (a), 1st paragraph, 7th line, after "grow" insert "to".
14 In footnotes, 1st line, delete "from", insert "form of".
36 No. 3, (b), 2nd paragraph, 14th line, delete "DBH", insert "dbh".
44 No. 3, (a), 2nd paragraph, 1st line, delete first "The", insert
44 No. 3, (a), 2nd paragraph, 4th line, before "largo" insert "last".
55 No. 4, (e), 2nd paragraph, last line, delete "that", insert "than"
53 No. 6, 1st paragraph, 5th line, delete "is", insert "are".
53 No. 6, 2nd paragraph, 5th line, after "area" delete ",".
59 No. 6, 4th paragraph on page, 3rd line, delete "patch", insert
RESOURCE MANAGEMENT REPORT
FISH AND WILDLIFE BRANCH
DEPARTMENT OF LANDS AND FORESTS
HON. RENE BRUNELLE
G. H. U. BAYLY
Digitized by the Internet Archive
RESOURCE MANAGEMENT REPORT
TABLE OF CONTENTS
No. 105 March 1970
EASTERN HEMLOCK ( Tsuga canadensis ): A REVIEW OF THE LITERATURE 1
- A. P. Matiece
1. Forest Geography 1
(a) Range 1
(b) Forest Associations 1
(c) Climate 1
(d) Physiography 2
(e) Successional Trends 2
2. Economics 3
(a) Markets 3
(b) Other Uses 4
3. Silvics 4
(a) Morphology 4
(b) Flowering and Fruiting 4
(c) Seed Production and Dissemination 5
(d) Germination, Seedling Development, Mortality 5
(e) Provenance 6
(f) Vegetative Reproduction 7
(g) Rooting Habits 7
(h) Growth Characteristics 7
4. Ecology 8
(a) Local Climate 8
(b) Light 9
(c) Moisture 9
(d) Soil 9
5. Artificial Reproduction 10
(a) Seed Collection, Extraction, Storage and Testing 10
(b) Planting 11
(c) Direct Seeding 11
6. Natural Reproduction 12
(a) Seedbed 12
(b) Light 12
(c) Cutting Methods 13
7. External Influences 14
(a) Response to Release 14
(b) Exposure 14
(c) Fertilization 15
External Influences (continued)
RUFFED GROUSE AND MICROCLIMATE
- H. R. Timmerman
A SURVEY OF WATERFOWL AND WATERFOWL HUNTING ON WALPOLE ISLAND INDIAN
- J. M. Collins
EASTERN HEMLOCK (Tsuga can adensis) :
A REVIEW OF THE LITERATURE
A. P. MATIECE, Deer Range Forester
Lake Simcoe District
1 - FOREST GEOGRAPHY
Hemlock occurs in southern Canada, the northeast
U.S.A., the Lake States, and throughout the Appalachian Mountains
to Georgia. In the northern part of its range it is found at
altitudes from sea level to 2,500 feet; in the south it is not
usually found below o00 feet (48).
In Canada, hemlock is found throughout the Mari-
time Provinces, in southwe stern Quebec, and in southern Ontario
as far west as Lake Superior (4).
(k ) Fores t As sociat ions
Hemlock is a constant element in the forest of the
northern hardwood region, occurring nixed with hardwoods, particu-
larly sugar maple, beech and yellow birch. It is also found in
nure patches of various size. In the northern part of its range
hemlock may be found with sugar mar>le, beech, yellow birch, white
ash, red spruce, balsam fir, red maole and red oak. In the south
it is usually found with shagbark hickory, tulip tree, basswood,
and various oaks (5, 48, 79).
The climate which favours hemlock and determines its
commercial range is predominately humid and cool (53, 93). Ex-
aminations of pollen grains in bogs in the southern U.S. indicates
that hemlock was once found considerably farther south and west
of its present range (16, 92). After the last period of glaciation,
about 11,000 or 12,000 years ago, a series of major climatic
changes brought about corresponding changes in forest vegetation.
After the ice receded a cold climate featured fir-spruce forests
which was followed by a cool dry period that favoured pines as
the major species. This was followed by a cool, moist climate
favouring hemlock and hardwoods, and a warm dry period featuring
oak-hickory forests (12, 91, 92). In Sanada
the V7arra, dry oak-hickory period did not occur.
Within its commercial range hemlock grows on a
wide range of soil conditions. It is more abundant and thrives
better under cool and moist, rather dry conditions (34, 48, 52,
97). Near the edge of its range its tolerance of soil and site
conditions becomes more critical (37, 38, 53, 57).
In the 1^7 Jihgland states Merril and Hawley (76)
claim that stand history ^cutting, fire), more than the soil is
likely to exert an influence on the presence or absence of
hemlock. It is unlikely that the situation in southern Ontario
differs in this respect.
(e ) Succassiona l Trends
Throughout its commercial range hemlock is accepted
as being a climax species, either by itself or in association with
beech or sugar maple. After a major disturbance the forest consists
first of pioneer species such as pincherry, aspen, white birch,
then pre-climax species such as white nine, red maple, red oak,
red spruce, and finally the climax species of hemlock, beech and
sugar maple (20, 65, 77, 97, 106).
Hemlock comes in under hardwoods and because of the
high tolerance to shade and its longevity and size, it can endure
and gradually penetrate the canopy and become the dominant species.
If a hemlock under story is present and the overhead canopy is
disturbed by cutting, windstorm, or disease, the young hemlock will
become dominant and the successionai trend will be speeded up. In
Nex* England, Pearson (o7) reports that hemlock spreads by seeding in
under hardwoods adjacent to existing hemlock stands and develops in
a concentric fashion around a central core of older trees. Martin
(72) indicates that in undisturbed conditions in Algonquin Park,
hemlock is slowly spreading throughout the hardwood forest. Bard (9)
concludes that if no fires or other major disturbances were to
take place, hemlock would continue as the dominant species since
it is self-perpetuating. At the edges oc its range hemlock is not
the climax species. For example, in Quebec where the boreal forest
meets the Great Lakes - 3t. Lawrence forest, Dansereau (25) reports
that hemlock is sub-climax to the deciduous forest.
Hemlock bark contains between 10 and 13% tannin
and in the past was an important source of tanning material.
Between 1850 and 1900 huge quantities of hemlock were cut for
bark to supply numerous tanneries both in Canada and the U.S.
Tanbark production figures from 1871 to 1901 as found in The
Canada Year Book 1905 (1) show an increase, a leveling off
and a decrease as the industry passed through its most active
phase (Table 1).
TANBARK PRODUCTION IN CORDS
YEAR CANADA ONTARIO
1891 329,797 110,111
1901 100,712 52,942
Siracoo County was the mam producer of tanbark
in southern Ontario, producing as much as nine adjacent counties
(6). In the U.S. tanbark production dropped from 1,170,131
cords in 1900 to 799,755 cords in 1905 and to 698,365 cords in
1909 (40). Pennsylvania was the largest producer with Wisconsin,
Michigan, New York and TJest Virginia following in respective
order. The value of hemlock lumber during this period was so
low that very rarely was any part of the tree other than the
More recently the market for hemlock lumber and
pulp^'ocd has improved. The wood is inclined to be somewhat
splintery and cross-grained, and is used more for rough con-
struction work than in a dressed condition. Typical uses are
for structural timbers, bridge planks, railway ties (treated),
boxes and crates, general construction and "or pulp (74).
* Sales of Hemlock bark, though nominally b}r the cord were actually
b3/ the ton, and in most cases the cord must x^eigh 2,240 lb. One
cord of dry, curled up bark might measure 4' x 4 f x 12' instead
of 4' x 4' x 8 1 , the standard cord sise. (40)
(b) Other Uses
Hemlock rates high in aesthetic value. It
is highly favoured as an ornamental, particularly in U.S.A.
and Europe (90). It is able to withstand the impacts of
recreational use (camping and picnicing) better than most
species as gauged by susceptability to disease, insect in-
fection and decline in vigour (88). It makes an excellent
hedge and is sometimes used for Christmas trees. It is an
important species in deer range, particularly in late
winter providing both food and cover for wildlife (50, 53).
The needles of hemlock tops felled in logging are eaten by
deer. Measurements by Stoeckeler et al (104) in the northern
hardwood -hemlock forest type show that hemlock will produce
117 pounds of fresh browse per square foot of basal area,
as compared with 13 pounds for hardwoods. Browse production
per M FBM (gross) of sawlogs cut is 1,507 pounds for hemlock
as compared with 121 pounds for hardwoods.
Hemlock is a long lived, mediism sized tree reaching
a height of 60-70 feet, and a diameter of 2 - 3 feet. The record
for age is 988 years for d.b.h. 34 inches, and for total height
160 feet (2). Onen growth trees develop dense pyramidal
crox-Tns with the lower branches touching the ground. The terminal
shoot is flexible and tends to curve away from the direction of
the prevailing wind. Self pruning is very poor. Dead branches
remain on the tree for many years (40).
& ) F lowering and Fru iting
The fruiting stage begins when the tree is between
20 and 50 years old and 15 to 25 feet high and continues until the
tree is over 400 years old. Suppressed trees under dense canopy
do not fruit regardless of age (3, 17, 53).
Separate male and female "lowers are found on the
same tree and appear in May or early June. Staminate flowers appear
before pistillate flowers (90). Pollination is by wind and maxi-
mum pollen receptivity takes place about a week before the lower
bud scales of the conelet turn back (01). After the pollen reaches
the female flower a period of about si:: weeks elapses before
fertilization takes place (81, 05). Cones mature by September or
October but are still green, wet and oily. The colour of the cone
changes from green to tan to brown and it then begins to open.
Hough (53) reports that only the central part of the cone
( c ) Seed Production jmd ^Dis s emina t ion
Hemlock is well knox-7n as a prolific seeder,
producing abundant seed at 2 - 3 year intervals with light
crops in intervening years. Seed dissemination takes place in
fall and winter. The hygroscopic cone scales open when dry,
close when x^et (2, 17, 19) and most of the seed is shed during
the first few periods of dry windy weather. This favours wide
seed dispersal. Olson et al (84), working in Connecticut,
state that much of the wintershed seed is sterile.
In good seed years huge quantities of hemlock seed
are produced. Lutz and Cline (66) in the Harvard Forest,
calculated that 11,700,000 sound seeds per acre were produced,
while Davis and Hart (29) at the Northeastern Forest Experiment
station, calculated that 13 1/2 million sound seeds per acre
were produced. This is considerably more than needed to ad-
equately seed the area. Hemlock seeds are relatively small,
about 200,000 seeds per pound (3, 48, 53). The cones persist on
the tree, after seed dispersal, until the following year.
(d) Germi nation, Seedling Development, Mor tality
Stratification (cold, moist storage) is probably
the most important factor affecting germination. Olson et al. (84)
and Heit and Eliason (49) say that new seed is incapable of
rapid germination in the fall when it is shed. Without stratifi-
cation average germination is about 20 - 30%. Baldwin (7) found
that germination increased from 7 to 45% with 30 - day stratifica-
tion and from 32 to 59% with 45 - day stratification. Further
tests by Baldwin (C) confirmed this, showing stratification at
39° to 50° F for 1-2 months averaged twice as high germination
after 30 days, three times after 60 da3^s and nearly four times
after 90 days than untreated seed. Under natural conditions, cold
moist stratification takes place during the winter and the seed is
ready for quick germination in the spring. Moist soil conditions
are necessary for rapid development of a root system and initial
survival of the newly germinated seedlings. If, due to lack of
stratification, incomplete stratification, or other causes,
germination is delayed until late in spring, the probability of
survival is greatly reduced (84) . Many of the newly germinated
seedlings will die during the summer when the soil becomes drier.
A wet summer resulting in a moist floor will ensure good survival
of hemlock seedlings. On the other hand, dry summers will
result in complete mortality of hemlock seedlings. Friesner
and' Potzger (39) in Indiana say that mortality of hemlock
seedlings is greatest in the first year and tapers off gradually
from there. Each added year of age, other things being equal,
gives the hemlock seedlings a better chance of survival. Davis
and Hart (29) at the Northeastern Forest Experiment Station
found that under normal circumstances, about 37 Q of the original
germinants will still be living after four years.
Shade plays an important part in the initial
survival and early development of hemlock. Establishment is better
in the shade than in full sun because of the drying effect of the
sun. Also, full sunlight can cause dark coloured seedbeds to
reach temperatures as high as 165° F, lethal to young seedlings
(53). Because Hemlock is a shade tolerant tree it can survive
and grow in the shade o: o--her trees for long periods of time.
Grasovsky (46) reports that if root competition were not present,
hemlock could survive under light conditions considerably lower
than that found in natural forest conditions. In New England,
Lutz (65) found hemlock regeneration was abundant in small open-
ings where partial or full shade is provided during the hottest
part of the day. Onl}' the south side of the strip (the shade
side) had hemlock regeneration on cut strips. Martin (72) says
that hemlock in Algonquin Park will regenerate where mineral
soil is not covered with a deep layer of litter.
Koroleff (59) after consulting with a number of
foresters, both from Canada and the U.S., concludes that hard-
wood leaf litter is detrimental to natural regeneration of hemlock.
Seeds may fail to germinate or germinate but fail on top of,
between or under fallen leaves. Small seedlings may be damaged
by mechanical act5.on of leaves (blown by xvind), or small seed -
lings may be killed by leaves which pile on top of them. A
single leaf falling on top of a cotyledon is sufficient to kill
it. Mortality from leaves is particularly bad in furrows where
leaves tend to collect and up to 100% of first year seedlings
may be smothered by heavy lea 1 : fall. Other studies by Davis and
Hart (29) and by Hough (51) confirm that hardwood leaf litter is
an important cause o p mortality of first year seedlings .
(e) Provenanc e
Observations by Nienstaldt, Olson, and others (CO,
02, S3, 84, 85) in the New England States and the Lake States in-
dicate that seedlings grown from southern seed source will grow
for a longer period of time than will those from northern seed
sources. This results in more growth per season for the south-
ern seeds. However, the southern seedlings do not harden off
soon enough in the fall and are susceptible to frost damage.
Similarly, mountain types tend to go dormant earlier than nearby
lowland types. Because of this, seed should come from areas with
a climate like that prevailing where the trees are to be grown.
(f) Vegetative Re production
Hemlock does not sprout naturally but may be
propagated by cuttings. Doran (30) reports that rooting is im-
proved by treatment with root- indue ing substances (indolebutyric
acid, Hormondin No. 3). Rooting of one year old twigs is better
than two year old or three year old twigs. These techniques are
suitable for horticultural purposes but are not practical in
A form of vegetative reproduction has been ob-
served in cut-over areas in northern New Jersey (27, 28). Hem-
lock stumps were putting on annual rings even though no leaf or
branch growth was present. In these instances a root graft to
an adjacent living hemlock tree was found to be responsible.
(g) Rooting Habits
The root system of hemlock is generally shallow
and spreading. This results in it being susceptible to damage by
surface fires, exposure, and drying out of the upper layers of
the soil. This type of root system, hox-jever, allows the tree to
grow on shallow, rocky soil where deep rooted species could not
1 ) Growth Characteris tics
The growth rate of hemlock in its natural condition
is ordinarily slow. For example, Morey (73) sampled seedlings and
advance growth under natural conditions in northwestern Pennsylvania
and found the following:
Heigh t ( feet) Age
Uith enough light growth can be rapid however. Vigorous seedlings
and saplings in light to medium shade nay grow 8-12 inches ?si
height per year, and in full sun with ample soil moisture, the
growth may be 18 inches or more (2) . Height growth begins early
in the spring (coincident with the start of needle growth) but
the rate is less than for ether species. However, the growth
continues cor a long period of time (85 to 95 days in New York
and Massachusetts) and thus outgrows many other species (21, 61,
62). Kozlowski (60, 61) says that height growth is closely
related to moisture conditions in the year preceding as a result
of the shoots being morphologically determined in the buds laid
down that year.
Diameter growth initiation in early spring may be
confused with rehydration of stem tissues. Considerable shrinking
and swelling in diameter takes place with changes in moisture. In
midwinter hemlock will decrease in diameter about 50% of the radial
increase from summer growth (60, 61, 107, 103).
Diameter increment of hemlock begins in the spring
earlier than for associated hardwoods. In the Chalk River, Ontario
area, diameter increment begins about the middle of Kay and by the
first week in August is practically completed (10) . Diameter in-
crement begins earlier, increases more rapidly, and lasts longer
for hemlock than for hardwoods. Frase-r (35) reports that the actual
time of commencement of radial growth is controlled by the late,
winter and early spring temperatures. Lack of moisture in mid-
summer will slow down or stop cambj.al growth. Measurements in
Wisconsin by Winget and Kozlowski (103) indicate that suppressed
trees are more sensitive to changes in environment than are dominant
Because individual trees may have been suppressed
for periods of time up to 200 years, any correlation between diameter
and age is unreliable. Gates and Nichols (41) state that most
mature hemlock stands have a good representation of trees of all
ages and so these hemlock stands could be considered all aged.
(a) Local Climate
.ock prefers cool, moist conditions. It is
usually more abundant on cool north e:nosures, steep banks of
river gorges, sides of cool ravines, swamp borders, and through-
out upland forests of white pine and hardwoods where drought
conditions are not present. Hemlock in pure stands produces an
e;;tremely dense canopy. As a result, the microclimate beneath
hemlock differs from the regional or open-station climate and
7rom that of adjacent hardwood stands, esnecially during the
growing season. Measurements have shown that this microclimate
is cooler and there is less evaporation during the growing
Light 5.ntensity under Dure hemlock stands is
reduced to 5 - 6% of open light intensity. Hemlock seedlings
are able to survive these conditions if soil moisture and
nutrients are sufficient. (Refer to 3 (d) this report)
In natural stands advance-growth hemlock often undergoes
periods of suppression ranging from 25 to over 200 years
(53). Under pure hemlock stands the ground cover is usually
formed exclusively of raw humus plants. Shrub vegetation and
seedlings of other species are usually absent.
Lyon (67) in New Hampshire and Marshall (70) in
Massachusetts report that a definite correlation between rainfall
and diameter increment for hemlock exists, particularly for hem-
lock which is groining on dry sites. Hemlock which grows on very
moist sites does not show this relationship. Diameter growth of
trees groining in moist conditions is greater than growth of trees
.growing on dry ridge conditions. In New England, Burns (14)
found that an increase in soil moisture results in an increase
of tolerance to shade and Bard (9) stated that higher moisture
(both atmosphere and soil moisture) favours the establishment of
hemlock regeneration. Drinkwater (32) found that light rains have
little effect on soil because much of the moisture is intercepted
by the dense canopy and evaporates before reaching the ground.
Hemlock stands produce a dry, acid litter which
decomposes very slowly (12, 32, 89). The nil of the upper soil
layers seldom exceeds 5.0 and is more commonly between 3.5 and
4.5 (12, 52, 53). This high acidity fosters podzalization
(leaching) of the upper soil layers. Even though the subsoil under
a hemlock stand and sn adjoining hardwood stand may be similar, the
surface soil under the hemlock will be more acid. Laboratory
experiments using seedlings in biotite showed that the pH changed
from 5.4 to 4.2 during the growth period of hemlock seedlings (98).
Because of dry loose litter under hemlock stands
the soil temperature fluctuates more readily with changes in air
temperature than under hardwood stands where the leaf layer acts
as an insulation (26, 36).
Due to the dense shade, dry, acid litter, and
shallow rooting creating severe root competition in the upper
soil layers, a young, fully -stocked stand of hemlock may exclude
any further reproduction of hemlock or any other wood or non-
woody vegetation for long periods (5, 55).
5 • ARTIFICIAL REPR O DUCTION
(a) Seed Colle ction, Extraction, Storage and Testing
Good seed years occur at 2 - 3 year intervals and
yields tend to be poor the first season after a good year. Cones
are most readily collected from the tops of trees felled in
normal cutting operations. There is a variation in time between
individual trees when cones change from green to tan. If cones
are picked green and dried quickly, they may turn pink or
purple rather than tan and fail to open unless subjected to
repeated cycles of re-wetting and drying at 100°F.
Cones collected just as they are turning brown will
open readily in the sun or in a dry room. They should be spread
out for a week or two and stirred occasionally so they will dr}?
quickly and not become contaminated by moulds . Green cones are
not only hard to open but are more subject to moulds.
Seed can be stored for 2-4 years in jars or
plastic bags a few degrees above freezing, but should be tested
periodically, as considerable variation in retention of viability
occurs. Storage at temperatures below freezing has also been
Seeds may be tested by cutting open with a razor
blade or finger nail to provide an estimate of the percentage of
filled, firm oily seeds with embryos. The percentage germination
will usually be less than this test indicates. If a more accu-
rate estimate is required, an actual germination test must be
carried out. Germination of hemlock is influenced by the inter-
action of photoperiod and temperatures, and optimum conditions
following stratification are 8 to 12 hours of light alternating
with 16 to 12 hours of total darkness, with corresponding alter-
nating temperatures of about 70° and 55 F respectively.
Continuous temperatures in the upper 70' s are definitely un-
favourable for hemlock germinating (84, 85).
Usually 3 year seedlings (3-0 stock) or 3 year
transplants (2-1 stock) are sufficiently well rooted and of
suitable size for out -planting. Poor survival of hemlock when
planted in the open is usually due to a lack of favourable
soil moisture conditions. Planting tests carried out at the
Northeastern Forest Experiment Station have shown that hemlock
will have good survival and growth rate if planted in orchard
type (mixed bush and openings) or under^lanted in a pole timber
stand (58). In the latter situation a thinning of the overstory
will be necessary to release the hemlock.
If seedlings are required for planting in the
spring earlier than the nursery is able to supply them, they
may be lifted from the nursery beds in the fall and stored over-
winter near the planting site. Tests by the Research Division,
Ontario Department of Lands and Forests have shown that ordinary
storage at 32°F or slightly less with a constant and high
relative humidity will enable the planting stock to be success-
fully carried over winter either loose or in crates. No elaborate
mechanism is needed and a simple insulated building is adequate
for storage (63).
( c ) Di rect Seeding
Seed should come from areas with a climate like
that prevailing where the trees are to be grown. If seed is
purchased, its age and source have a bearing on germination and
development ('see 3 (e) and 5 (a) this rer>ort). If seeding is
done in the spring, the seed must be stratified. Seed sown in
the fall will be naturally stratified and ready for quick germ-
ination in the spring. However, there nay be overwinter losses
to rodents, birds, fungi or other causes. Tests by Jordan and
Sharp (58) in Pennsylvania, suggest that seeding in the or>en or
under a closed canopy will probably give poor results; seeding
in orchard types (mixed bush and openings) will likely give
better results. In all cases a suitable seed bed as indicated
in section 6 (a) of this report is required.
6 . NATURAL REPRODUC TION
Dry litter seedbeds are unfavourable for germina-
tion of hemlock. Maissurow (69) contends that fire is necessary
for the natural regeneration of hemlock but this is questioned
by Graham (42) and others (see 1 (e) and 7 (h) this report).
Favourable seedbeds consist of moist, well decomposed litter,
rotted stumps and logs, mineral soil, moss, or even moss-
covered boulders. After germination, continuing moist
conditions are necessary for survival. Hough (51) in Pennsylvania
and Olson et al (35) in Connecticut report that naturally shaded
seedbeds are usually sufficiently moist in early spring for
initial survival but summer drought may eliminate many of the
seedlings. Mineral soil, although an excellent seedbed for
germination, subjects the young seedlings to washing-out or
mudding-under in heavy rains and to frost heaving. In addi-
tion, exposure to direct sunlight and the resultant high
temperatures can kill the young seedlings either directly or
through drying out the upper layer of soil. Germination of
hemlock on mineral soil can be very high. Davis and Hart (29)
report up to 3.8 million seedlings per acre, so even though
mortality might be high, sufficient numbers may remain. Mineral
soil seedbeds still remain receptive to hemlock four years
after being exposed.
A seedbed consisting of a mat of hardwood leaf
litter is unfavourable for the establishment of hemlock regenera-
tion. (see 3 (d) this report) Removing hardwood trees to reduce
the quantity of leaves falling on a newly germinated seedbed
would eliminate this hazard. On the other hand, a sufficient
number of overhead trees should be maintained to provide partial
shade on the seedbed.
In very dense shade conditions, such as under a
pure young hemlock stand where daylight intensity falls below 100
foot candles, hemlock will germinate but may subsequently be
killed or stunted. Bard (9) and Steams (99) report excellent
hemlock regeneration is obtained in small openings ( ap proximate ly
0.1 acre) created by natural conditions, e.g. windstorm, or
artificially, e.g. cutting. Lutz (65) reports that hemlock
will regenerate well on shaded south sides of cut strips in
In clear cutting operations, if hemlock is to
be maintained, it should be established in the understory be-
fore the overstory is removed.
(c ) Cutting M ethods
Clear cutting or making large openings in mixed
hemlock-hardwood stands will result in dense reproduction of
hardwood pioneer species, hardwood sprouts, and the virtual
exclusion of hemlock. The fact that hemlock does not sprout, as
opposed to hardwoods, and that large openings dry out the site,
makes this type of cutting unfavourable to regeneration of hen-
lock. Studies in Nova Scotia by Martin (71) also show that clear
cutting followed by fire is almost certain to eliminate hemlock.
Host forms of partial cutting will favour re-
generation of hemlock, provided that a seed source and a suitable
seedbed (see 6 (a) this report) is present. First cutting of
the uniform shelterwood system gives good results , as does group
selection and strip cuts, particularly if logging is carried out
in the summer so that accidental scarification takes 10 lace (2.4,
34, 65, 66, 70).
Luts and Cline (66) working on the Harvard Forest
renort that the openings should be at least 40 feet wide in
group selection cutting. Strips should be oriented in an east-
west direction since this provides more shade on the south side
of the strip. Considerable hardwood will probably come in on
the strips along with the hemlock.
Seed tree cuts leaving from 1 to 10 hemlock seed
trees per acres have given natural regeneration of hemlock in
southern New York (76). Hardwoods come in quite heavily as well
and if hemlock is wanted, it will be necessary to carry out clean-
Both Hosley and Ziebarth (50) and Spurr (97) report
that most of the hemlock stands in Massachusetts present today
originated either from selective cutting, which removed the hard-
wood or white pine overstory, freeing the hemlock understory,
or developed through persistance in areas which had never been
Logging slash of hemlock, which will persist for
up to 15 years, may interfere x^ith the immediate establishment of
regeneration if it is dense (24) but probably will have no notice-
able effect in the ultimate density of stocking (75).
7. EXTERNAL INFLUENCE!
(a ) Response to Release
Hemlock has the ability to survive for many }?ears
in a suppressed condition and then recover and grow at an ac-
celerated rate when it is released. Marshall (70) reports the
rate of growth after release was 5 1/2 times what it had been
before release. Martin (72) tells of one tree in Algonquin
Park which took 108 years to grow 3 inches d.b.h. under
suppression and only 80 additional years to grow 20 inches d.b.h.
after release. The response to release is influenced by
stand density. Well stocked stands will have greater accelera-
tion in growth than poorly stocked stands. Tor a given
diameter the older the tree when released, the greater its re-
sponse will be. Suppressed trees, after release, generally over-
take open grown trees (53, 70).
Results of partial cutting in the Lake States
show that only the well formed trees are able to respond to re-
lease. Low vigour trees will probably die when exposed to
direct sunlight (22). (See 7 (b) this report).
Pv.eleas.ing hemlock by spraying competing hardwoods
with Kuron" and 245 OS* showed little damage was done to the
hemlock by the chemicals, even when spray was applied so heavily
that it was dripping off the leaves. Spraying took place in
August when the new growth of conifers had hardened off. Only
a few terminal buds did not put out new growth the following
Hemlock is subject to post-logging death because
of exposure. This applies particularly to low-vigour trees. In
his studies in northern Michigan, Graham (43) found the amount of
increased exposure and the size and shape of the crowns of the
Kuron - a propionic ac5.d from 245T reputed to be more effective
against maple and oak than the normal acetic acid form
245 OS - 245T plus oil stable emulsifier
residuals have a bearing on susceptibility to exposure. Generally,
trees having heavy crowns of conical shape are able to withstand
exposure, and those having narrow, flat-topped crowns are not
able to withstand exposure. Exposure on the south and west is
more likely to cause death than exposure from other directions.
Hemlock is likely to die if sunlight falls directly on the trunk
and exposed roots.
When cutting, any groups of hemlock to be left
should be sheltered especially from the south and west by maple or
other trees that are resistant to exposure. All hemlock with
short or narrow crowns should either be marked for cutting or,
if reserved, left completely sheltered from the south and west
sides. Most hemlock with crowns at least 25 feet wide and
occupying 2/3 or more of the total height may be exposed without
much danger of injury, but if practicable, even these should be
given some protection from the south-west.
Burnham et al (13) in Pennsylvania and Lutz (65)
in southern New England recommend release of hemlock seedlings
or suppressed hemlock from hardwood overstory should be gradual
in order to minimise damage from exposure. Observations of
cutting operations in the northeastern United States indicate
that partial cutting or thinning of a hemlock, stand should be
light in order to hold hemlock mortality to a minimum (33, 64).
( c ) Fertilization
Although hemlock seedlings can survive at a low
level of nutrition, growth can be increased several- fold by main-
taining nutrition at moderate levels, llaintenance of high levels
of nutrition, or the continuance of fertilization and watering
into the late summer or fall should be avoided because the slants
may continue growth too late in the season and might not be
sufficiently hardened off before cold weather sets in. Over-
fertilization of large hemlock increases their susceptibility
to -^ bizoctonia , a root-destroying fungus. If the roots are not
too severely injured, the trees will recover if the soil is
treated with - hydro::yquinoline sulphate (84, 85).
(d) I nsects
Of more than 20 insects having hemlock as host,
only the hemlock looper, Lambdjoia f^scetl^Tia and the heralock
borer, Melansphila fulvog uttata are economically important.
The hemlock looper is a defoliator which reduces the abundance
of hemlock when it becomes prominent. Graham (43) indicates that
where the upper crown canopy is composed of 75% or more of hem-
lock, extremely dense modulations of hemlock loopers develop,
completely defoliating and killing all hemlock trees. When the
proportion of hemlock is low, the insects are few and cause
little or no injury. Hemlock borers attack the trees when they
are in a weakened condition, such as from drought. Graham (43)
states that these insects cannot complete their development in
Hemlock is surprisingly free from serious diseases.
Worthy of note are Forties pini , a trunk rot found in mature trees
an< ^ Armillaria roellea, a shoestring root fungus, x^hich attacks
trees when they are in a weakened condition (34, 43, 94). Young
seedlings are also subject to damping-off. windshake is common
in hemlock and the species is extremely susceptible to sulphur
(f ) Weather
Hemlock is seriously affected by drought conditions.
Heavy mortality has heen recorded particularly on drier sites,
ridge tops, southern exposures, shallow soils and steep slopes.
The shallow rooting system of hemlock makes it more susceptible
than most other species to drying out of the upper soil surface.
Secrest et al (94) found that during severe drought conditions
no annual rings are put on in the lower portions of the tree al-
though rings are put on in the upper portions of the tree. No
rings are put on for a 2 to 5 year period prior to the death of
the tree. Lutz and Cline (66) and St5,okel (101) report that
during dry summers there is a heavy loss of seedlings (see also
3 (d) and 6 (a) this report).
Hemlock is able to withstand damage from ice storms
extremely well (15, 31, 53, 56, 96). Greater resilience of the
branches, smaller upper crown, and branching habits better adapted
mechanically to resist weight on the crown probably account for
Winter browning can occur when warm weather along
with bright sunshine and drying winds come after the soil is
frozen. The foliage above snow line to a point 2 or 3 feet above
turns brown and defoliation may occur. Stoeckeler and Rudolf
(103) working from the Lake States Forest Experiment Stat?lon re-
port that recovery from this condition the following summer is
usually good. Trees with up to 41% defoliation suffered no mort-
Frost damage to seedlings is not common but
occurs occasionally on south facing clones. Loss from wind-
fall is not great except iu a wet situation.
Hemlock can be adverse!}?- affected by deer,
porcupines and varying bares. These animals eat the foliage and/
or bark of the tree. In addition, red squirrels, chipmunks, and
some birds eat the seed. Numerous references indicate that hem-
lock foliage is a preferred food of deer. Maynard et al (73)
reports that in cold weather (0° F and below) hemlock is eaten
in greater quantity than in warmer weather. Reports from Wis-
consin, Michigan, i enncylvania and Algonquin Park, Ontario state
that in areas with heavy deer populations, hemlock seedlings
are browsed close to the ground repeatedly year after year so
that they never have a chance of becoming established (12, 34,
44, 54, 100, 105). Brown and Curtis (12) in Wisconsin and
Graham (45) in Michigan warn that this could result in the
forest developing into almost pure hardwood with virtually no
winter cover. Hemlock seedlings have been completely eliminated
outside exclosures or have been seriously reduced in numbers in
some areas of the U.;j. Usually only the foliage and small twigs
of hemlock are eaten by deer but they have been reported eating the
bark of young hemlock trees in some areas of heavy populations
in Pennsylvania (18). Hhere deer browsing ores sure is high,
seedlings should be protected. Grises (47) found that
substantially larger numbers of stems of the commonly browsed
s-oecies were found in slash piles than in intervening openings.
The possibility for deliberately using slash to protect seed-
lings might be considered.
Hemlock is also a preferred winter food of the
porcupine (11, 23, 102). Much of the feeding is foliage brows ing
of larger trees and damage of this sort is difficult to appraise.
Damage can be severe where the feeding takes the form of gnawing
the bark, sometimes girdling the tree and killing it.
Varying hares also browse young hemlock but Ostram
(>_ G) reports that damage is usually considerably less than the
damage by deer.
Because of its shallow rooting system hemlock is
particularly susceptible to fire in spite of its relatively thick,
fire resistant bark. Ground fires- will destroy all hemlock re-
generation and a fire following a clear cut will result in
regeneration of pioneer species. Even a light fire may
practically eliminate hemlock. To ensure the continuation of
hemlock, Zon (109) states that complete fire protection is
C. HEMLOCK MANAGEMENT
Growing hemlock in Ontario within its natural
range should not prove to be a difficult problem provided that
the requirements of the species are recognised and provided for.
Since hemlock will do better on cool moist conditions than on
droughty ones, it would be advisable to concentrate management
on these conditions. To achieve natural regeneration, the
combination of (1) a seed source, (2) a suitable seedbed and
(3) adequate light must be provided, .jcarification, either
deliberate or accidental through logging activity in close
proximity to standing, seed-bearing trees will provide the first
t^?o requirements. A partial opening of the overhead canopy
whether uniformly or in patches will give sufficient light and
partial shade; a condition best suited to regeneration of
hemlock. Similar light conditions are desirable if hemlock is
to be planted.
When hemlock seedlings become established it is
possible that competition from hardwood seedlings may become
severe, necessitating a cleaning to release the hemlock. As the
overhead canopy closes in and light in the understory is reduced,
removal of a portion of the over story to release the hemlock
would be advisable.
Where heavy deer populations are present planting
of hemlock should either be avoided or the young seedlings should
be protected from, browsing. Fencing the entire planted area,
placing screening around individual seedlings, piling logging
slash around seedlings, or using repellants are all possible
methods of providing protection.
Tvhere natural regeneration or advance growth of
hemlock is already present in the understory a thinning or release
cutting of the overstory will hasten the development of the hem-
lock. Care must be taken, particularly on dry sites, that the
overstory is not opened so much as to e:mor>e the understory hem-
lock to total direct sunlight.
Periodic thinnings as the stands progress
through the polewood stage to maturity will insure that
growth will be maintained at a high level. Fertilization
may increase growth on sites which are deficient of nutrients.
After growing out of reach of browsing deer,
hemlock stands are generally not affected seriously by wildlife.
On the other hand hemlock stands provide valuable winter cover
As the stand grows to maturity it may provide a
number of useful functions; erosion control on steet> slopes or
shallow soil ridge tops, food and shelter for wildlife, pleasing
vistas before achieving its final use as a wood product.
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78. Morey, H. F. 1936. Age-size relationships of Hearts Content, a virgin forest
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81. Nienstaedt, H. and H. B. Kriebel. 1955. Controlled pollination of Eastern
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82. Nienstaedt, H. and J. S. Olson. 1961. Effects of photoperiod and source on
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96. Spaulding, P. and A. W. Bratton. 1946. Decay following glaze storm damage
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97. Spurr, S. H. 1956. Forest associations in the Harvard Forest. Ecol. Monog .
98. Spyridakis, D. E., G. Chesters and S. A. Wilde. 1967. Kaolinization of
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99. Stearns, F. W. 1949. Ninety years change in a Northern Hardwood forest in
Wisconsin. Ecology 30: 350-358.
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101. Stickel, P. W. 1933. Drought injury in hemlock-hardwood stands in Connecticut,
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The following publications were also used as source material although no
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Arbogast, C, Jr. and M. L. Heinselman. 1950. Damage to natural reproduc-
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Bourdeau, P. F. and M. L. Laverick. 1958. Tolerance and photosynthetic
adaptability to light intensity in White Pine, Red Pine, Hemlock and
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Bromley, S. W. 1935. The original forest types of Southern New England.
Ecol. Monog. 5: 61-89.
Buell, M. F., et al. 1966. The Upland Forest continuum in Northern New
Jersey. Ecology 47: 416-432.
Burns, G. P. 1923. Studies in tolerance of New England trees. LV minimum
light requirements referred to a definite standard. Vt . Agric . Exp.
Sta. Bull. 235.
Daubenmire, R. F. 1930. The relation of certain ecological factors to the
inhibition of forest floor herbs under hemlock. Butler Univ. Bot. Studies
Ehrhart, E. 0. 1936. Forest management and deer requirements on the
Allegheny National Forest. Jour. For. 34: 472-474.
Friesner, R. C. and J. E. Potzger. 1936. Soil moisture and the nature of
the Tsuga and Tsuga-Pinus Forest associations in Indiana. Butler Univ.
Bot. Stud. 3: 207-209.
Frontz, L. 1930. Deer damage to forest trees in Pennsylvania. Penn. Dept .
For. & Waters, Res. Circ. 3.
Frothingham, E. H. 1931. Timber growing and logging practice in the Southern
Appalachian region. U.S.D.A. Tech. Bull. 250.
Frothingham, E. H. 1943. Some observations on cutover forests in the Southern
Appalachians. Jour. For. 41: 496-504.
Gilbert, A. M., R. W. Wilson, Jr. and R. J. Hutnik. 1955. Growth behaviour of
Northern Hardwoods after a partial cutting. Jour. For. 53: 488-492.
Graham, S. A. 1941. Climax forests of the Upper Peninsula of Michigan.
Ecology 22: 355-362.
Harper, R. M. 1952. Hemlock in Alabama: a supplementary note. Ecology 33:
Hawes, A. F. 1912. Hemlock in Vermont. Comparative study of log rules. Vt .
Agric. Exp. Sta. Bull. 161.
Hawes, A. F. 1923. New England forests in retrospect. Jour. For. 21: 209-
Hosley, N. W., et al. 1931. Wild animal damage to New England forests. Jour.
For. 29: 700-708.
Hough, A. F. 1936. The dying of hemlock and other species on the Allegheny
National Forest. Allegheny For. Exp. Sta. Tech. Note 9.
Jones, E. W. 1945. The structure and reproduction of the virgin forest of
the North Temperate Zone. The New Phytologist 44: 130-148.
Kilburn, P. D. 1960. Effects of logging and fire on xerophytic forests in
Northern Michigan. Bull. Torrey Bot . Cl. No. 87: 402-405.
Longwood, F. R. and W. A. Salminen. 1952. Reproduction after cutting in
hardwood -hemlock stands. Lk. States For. Exp. Sta. Tech. Note 379.
Oosting, H. J. and D. W. Hess. 1956. Microclimate and a relic stand of Tsuga
canadensis in the Lower Piedmont of North Carolina. Ecology 37: 28-39.
Pearce, J. 1937. The effect of deer browsing on certain Western Adirondack
Forest types. N. Y. State College of Forestry Bull. 10.
Piussi, P. 1966. Some characteristics of a second-growth northern hardwood
stand. Ecology 47: 860-864.
Putham, D. F. and L. J. Chapman. 1938. The climate of Southern Ontario.
Scientific Agriculture 18: 401-446.
Scholtz, H. F. 1930. How long does hardwood slash remain a fire menace?
Jour. For. 28: 568.
Segars, C. B., L. C. Crawford and A. M. Harvill. 1951. The occurrence and
distribution of hemlock in Alabama. Ecology 32: 149-151.
Spurr, S. H. 1956. Natural restocking of forests following the 1938 hurri-
cane in Central New England. Ecology 37: 443-451.
Stearns, F. W. 1951. The composition of the Sugar Maple-Hemlock-Yellow Birch
Association in Northern Wisconsin. Ecology 32: 245-265.
Stegeman, L. C. 1937. A food study of the white-tailed deer. North Amer.
Wildlife Conf. Trans 2: 438-445.
Stephenson, A. B. 1959. Deer investigations in Biggar and Wilkes Townships,
Algonquin Park. Res. Div., Ont. Dept . Lands & Forests, Sect. Rept . (Wildlife)
Stoeckeler, J. H., R. 0. Strothmann and L. W. Krefting. 1957. Effect of deer
browsing on reproduction in the Northern Hardwood -Hemlock type in
Northeastern Wisconsin. Jour. Wildlife Mgt . 21: 75-80.
Watson, R. 1925. Notes on natural regulation and growth of Northern Hemlock
and hardwood forests. Jour. For. 23: 936-940.
Westveld, R. H. 1933. The relation of certain soil characteristics to forest
growth and composition in the Northern Hardwood Forest of Northern Michigan.
Mich. State Coll. Agric . Exp. Sta . Tech. Bull. 135.
Zon, R. and H. S. Graves. 1911. Light in relation to tree growth. U.S.D.A.
For. Serv. Bull. 92.
Zon, R. and R. N. Cunningham. 1931. Logging slash and forest protection.
Wise. Agric. Exp. Sta. Res. Bull. 109.
RUFFED GROUSE AND MICROCLIMATE
H.R. TIMMERMAN, Biologist
Thunder Bay District
/in under standing of the basic ecology of Ruffed
Grouse, (B onasa umbellus L.) is essential to its management.
Ruffed Grouse habitat has been studied extensively with respect
to plant species and plant densities, Edrainster (1947). Little
if any work has been conducted on the less obvious environmental
factors such as microclimate.
Ruffed grouse select various cove:: tynes at
different times of year. The differences in microclimatic
aspects of temperature, wind velocity, and solar radiation have
a significant effect on cover tyoe selection. Dennis (1^66)
conducted a study on the selection of cover types by Ru'fed
Grouse in Wellington County, Ontario, Much of the information
collected during this study will be referred to in the following
discussion dealing with grouse and microclimate.
1 • GENERAL .BACKG ROUND INFORMATION
Bump, Darrow, Sdminster and Orissey (1947) devote
915 pages and 16 years of investigational work in their monumental
text, ''The Ruffed Grouse Life History, Lrcpagation, Management"'.
The following generalities are the only clue given to describe
the types of microclimate preferred by grouse.
(a ) Brood Cover Ch oice
Temperature, wind and atmospheric conditions might
logically be expected to affect brood cover choice. There is a
tendency for grouse broods to seek the mere ooen cover of hardwoods
during cold periods and of conifers during warm weather. Overgrown
land types and second growth hardwoods were found to be most
frequented when temperature conditions were colder than normal.
Young hardwoods and conifers and conifers alone were patronized
more on excessively warm days. The deep, often moist, cool
depths of older patches are also o : ton "roquented. Mature grouse
are inclined to be ill o.t ease on windy days. Youngsters too
normally move around less on such days. Grouse broods prefer
some type of cover during rainy weather and the more onen over-
grown slash areas arc less well patronized at this time. The
reverse tends to occur during sunny and cloudy weather.
( b ) Adult Coyer Choice
Birds are inclined to novo into mixed second
growth hardwoods and conifers or thick conifers, on colder than
normal days. Conversely, on warmer than normal days, adult grouse
tend to move to some o r the more ooen r.ovev types deficient o r
conifers. During windy weather, thick coniferous cover anpears
to be less frequented. Uind normally seems to exert a minor
influence in grouse cover choice. During snow storms the birds,
as expected, are most likely to seek out heavy coniferous cover.
However, no cover is completely shunned even in snowy weather.
Rain seems to bother grouse less than snow. Fewer birds seek
coniferous cover and more seek mixed second growth and conifer-
ous stands .
Ground conditions exercise a greater effect on
grouse cover choice than is generally realized. There is con-
siderable variation in type use, deoending on whether the ground
is dry, wet or snow covered.
( c ) Nesting covers
Second growth hardx^oods arc predominantly used as
spring nesting grounds. The presence of considerable summer :'ood
and the more open character of under gr ox 7th and ground cover in
this type, in contrast with that exist ing in most overgrown
lands and slashings, may furnish a clue.
Generally grouse choose the warmest sites in winter,
the warmer roosting sites at night in summer and the coolest spot
:o rest during a warm summer day, Hungerford (1951). In Idaho,
Hungerford found greater importance in grouse use of various
microclimatic zones than in cover tyres. This is due to the high
elevations which range 7rom 3,000 to 3,500 r eet. Local differen-
ces in climate are well recognized in sore fields but their effects
on wildlife behaviour have scarcely been suggested. It is known
that there &:ce great differences in microclimate between Forested
and non- forested areas. It is also known that a forest clearing
has a different microclimate :rom that of either the unbroken
forest or non- forested land. It may well be, says Hungerford,
that the microclimatic differences are o r>rime importance in
raking the edges of clearings or o" cover types so attractive to
wildlife. In Idaho, because of the mountainous terrain and the
inversion phenomenon, the. ridges offer a warmer habitat in both
winter and summer and night and day.
2 • GROUSE AND MIC ROCLU ftTE IN THE S T. LAv /RENCE FOREST REGION
Under suitable snow conditions, grouse tracks and
roost sites are useful as indicators of the relative use that
cover types producing different microclimatic conditions receive.
The study carried on by Dennis (I960) examined microclimatic con-
ditions in w inter and spring.
(a) Cover type s
Dennis ( 1966 ) conducted his studies on an area of
125 hectares. The area was surveyed and a vegetation map prepared
A total of 11 major cover types were established. The greater
proportion of the study was conducted on two of these cover ty^es
which are referred to in this naper as (a) and (b).
Cover t ype (a) Area 8.15 hectares
i Most com mon species:
.5 - 2.5 metres: white cedar, honeysuckle, elder-
berry, nightshade, red osier,
2.5-6 metres: scotch pine, white pine, x?hite
spruce, norway spruce
6 -:- metres: scotch pine, white pine, norway
The tree canopy cover in. this oredominantly coni-
ferous stand averaged 50%. Ground cover was moderate to heavy in
all seasons of the year. Much white cedar regeneration had
occured, and elderberry and nightshade were present in the more
C over t ype (b) Area 4.27 hectares
ii Most common soecies:
.5 - 2.5 metres: red osier, dogwood, choke-cherry
2.5-6 metres: trembling asr>en, white pine, white
spruce, black cherry
6-!- metres: white elm, white cedar, balsam-
poplar, trembling aspen.
This bottomland cover type was very open with
only 15% canopy. Ground cover was heavy during the summer but
more open in winter.
Thus it can readily be seen that the basic cover
type is coniferous and deciduous.
3. MICROCLIMATE CHA RAC TERISTICS OF (a) CONIFEROUS COVER A ND (b )
DECIDUOUS COVER "
Both Geiger ( 1965 ) and Upurr ( 1957 ) state that
coniferous cover moderates temperature extremes. Conifers prevent
a high percentage of solar radiation from reaching and heating the
air mass beneath them, thus reducing daily maxima. Geiger goes on
to say that at night a coniferous cover produces effects similar
to those of clouds in an open area. These effects are caused by
absorption of the radiation from the snow by the crown cover and
reradiation to the snow surface. This reduces heat loss within
the stand and results in a higher minimum temperature. Kittridge
( 1943 ) states that evergreens are more effective in winter in
reducing maximum and raising minimum air temperatures. Daily
temperature extremes are much less in upland conifer cover types.
A bottomland conifer cover usually has lower mean minimum, and
maximum temperatures than an upland site. Here the effects of
topography may be greater than vegetation.
Dennis ( 1966 ) found that during January, February
and March, the lowest density conifer stands produced the greatest
daily temperature extremes. These are also areas containing a high
percentage of deciduous cover. Deciduous stands oroduce more air
mixing and hence greater daily temperature fluctuations (Geiger
1965). The effectiveness of a forest canopy in reducing the in-
tensity o" solar radiation increases with increase in (a) percent-
age of crown cover (b) closeness of spacing (c) tolerance of species
and (d) progression of natural succession towards climax ( Kittredge
Dennis recorded temperatures in both cover tyoes
(a) and (b). The highest mean maximum in both cover types occured
at a height of 3 metres while the lowest mean minimum occured at
15 centimetres, which approximates grouse height. Sparks and Buell
( 1955 ) found a lower minimum at 20 centimetres than at 2 metres
over snow. This lower minimum at grouse height ( 15 cm. ) may
be partially due to the fact that snow radiates strongly at night
and hence temperatures are lower nearer the snow ( Geiger 1965 ).
Spurr ( 1957 ) found a similar situation in the Harvard forest.
Figure I illustrates the mean minimum and maximum temperatures
recorded under (a) and (b) cover types.
A higher wind velocity occurs in a typical open
deciduous cover type than in a coniferous cover type. Also
higher x*ind velocity means increased dynamic convection resulting
in a lower temperature near the ground during the day ( Geiger
1965 ) . Both Geiger and Kittredge ( 1948 ) state that as wind
velocity increases, the percentage penetrating the forest de-
creases. Velocities in the forest are characteristically low,
usually ranging from 1 to 2 m.p.h. on the average. Dennis ( 1966 )
found that with grouse at least two factors affect wind velocity.
The first is cover density and the second is that there is a
dependence on the position of the cover with respect to open areas.
Sjx open field along a forest edge tends to increase wind velocity.
Dennis recorded wind velocity for 322 grouse flushes over a two-
year period. Of these, 310 sites had a wind velocity of less than
1 netre per second. Two flushes were recorded with velocities
between 1 and 2 metres per second and 2 flushes between 2 and 4
metres per second.
Grouse frequently day roost in areas where sola"
radiation is unobstructed. i)ennis simulated a grouse by building
a model which could duplicate average internal core temperatures .
He found that the use of these day roosting areas, may have re-
duced heat loss on surrrrj/ winter days. This reduction is caused by
reflected solar radiation and insolating qualities of snow warming
the surface of the bird. The warmed surface increases the ambient
temperature and a decreased gradient between core and ambient
temperature results. The capabilitjr of the surrounding air to act
as a heat sink is reduced thus reducing heat loss by 50% during
sunny days in late winter. During April, Dennis ( 1966 ) recorded
a heat loss in a 15% canopy open cover type which was 537 less
than the heat loss in a 95% conifer canopy.
(a ) C over type variable s and buffed Grouse Utilization
Utilization of cover tyoos by ruf <:ed grouse appear to
depend on food, shelter and climatic factors within the cover tvnes
( Dennis 1966 ).
(b) W inte r loafing, feeding and roosting covers
Marshall ( 1965 ) believed that the paramount factors
affecting ruffed grouse behaviour during cold periods involved a
quest for habitat that would minimize energy loss and for foods of
MEAN MIN. AND MAX. TEMPERATURES RECORDED UNDER
A--UPLAND CONIFER AND B--BOTTOMLAND HARDWOOD (DENNIS. 1966)
1 1 1 1
Mar. 2-7 Mar. 8-13 Mar. 14-20 Mar. 23-29 Mar. 30-Apr. 5 <-
A-3 metres — i
mean max .
types A & B
4 A-5 metres— »
B-15 cm. _
mean mm .
types A & B
high caloric value. Hunger ford ( 1951 ) celt that many forms of
cover were used only because they were situated within the most
suitable microclimatic zone. During the day mean maximum tempera-
tures are higher in a more open canopy cover type. Hence solar
radiation reduces grouse body heat loss to a greater extent in
such an area than in a more sheltered cover type. Dennis found
that solar radiation was able to penetrate the dense choke-cherry
stands which provided protection from avian predators.
During the winter most of the feeding is done on
t3:ees. Aspen buds are a favourite grouse food at this time of year.
Marshall ( 1965 ) indicated that flower buds of staminate aspen
trees were a very rich source of fats and protiens. Cover types
containing a good supply of mature aspen trees will be heavily
used by grouse in January and February ( Dennis 1966 ) . Grouse
may also supplement this supply with other foods such as night-
shade berries and ash keys. Marshall also indicated that juvenile
male grouse in Minnesota moved a limited amount duri.ng the winter.
Movements appeared to have been limited to leaving roost sites to
feed on aspen buds and returning to day or night roosts, depending
on the feeding period. "For actual roosting the younger conifer
trees are chosen. Clumps of 3 or more trees from 2-6 inches
DBH comprise a typical roosting site. The roosting perch is
usually 8 - 10 feet above the ground. Dorney ( 1960 ) found
that lightly crusted snow that grouse cannot penetrate, in cold
weather can again become "soft" allowing birds to burrow under
it. Exposure, cloud cover and wind as well as temperature all
affect the crusting of snow. Above average winter temperatures
may, therefore, be unfavourable to grouse.
Snow roosting normally occurs during severe winter
weather Edminster (1947). True snow roosting occurs when the
grouse is completely buried. Partial snow roosting may also occur
in both day and night roosting areas. The following table in part
prepared by Dennis ( 1966 ) illustrates that snow roosting in cold
weather would reduce heat loss when compared to tree roosting.
As shown in Table I temperature ranges under the
snow were much reduced from those above. As snow depth increases,
temperatures under snow will be very close to freezing even
though the air temperature may be as low as - 33°C (Geiger 19.65).
The effect of snow depth was illustrated by the minima occuring
during the periods January 3 - February 5, and February 19 to
February 24. The mean above snow minima differed by only 0.2 ( ' C
( -16.4 and -16.2°C ). During the same period the difference
between the under snow temperature ( -4.0°C )and the above snow
temperature ( -16.4 C ) was - 12.4°C while in the period February
19 - 24 the difference was -9.2°C Grange (1949) feels that
when snow roosting becomes impossible because of crusting,
grouse become subjected to an increased rate of oredation.
With hard snow, predators may hunt with greater intensity
because the crust provides protection from grouse buffer
species such as rodents. Snow roosts tend to make grouse
vulnerable to fox predation. It appears foxes learn to look
for breather holes which are always present when grouse are
completely snow covered. Grouse often resort to trees,
especially conifers for roosting, or when disturbed by a
predator or intruder.
TABLE I - MEAN MINIMUM AND MAXIMUM TEMPERATURES OBTAINED FROM
RECORDING THERMOMETERS - DENNIS (1966) IN COVER
TYPE (a) UPLAND CONIFER
Period Air temp 15 cm above snow
, mean mi n °C mean max.
Jan. 23 - 29 -11.6 -4.7
Jan. 30. Feb. 5 -16.4
Feb. 6-12 - 3.C
F-b. 13 - 18 -12.2
Feb, 19 - 24 -16.2
denth Snow temp under snow
, . cm mean mla °C mga n max .
10' -3.9 -4.7
(c) S pring dr umming 3 feeding and nesting c overs
In southern Ontario a shift in utilization o. r cover
types usually takes place in late March or early Anril (Dennis 1966).
At this time male grouse move to their spring drumming territories
and a general movement from wintering habitat occurs. Palmer,
(1961) found that ground vegetation Ttfas less dense and large- shrub
and tree cover more dense in drumming site locations than in
surrounding habitat. He believed that a combination of variables
rather than vegetation density alone, governed the choice of
drumming areas. Dennis found that the majority of drumming sites
occured in cover having the lowest mean wind velocities and the
highest minimum temperatures during the spring. All drumming sites
observed were located under at least 60% canopy cover.
Open cover types become ba.-:c of snow first in the sr>ring
These areas usually produce the first succulent green
plants on which grouse -reed extensively. Cover at the edge
of these open spring feeding areas provides shelter and escape
cover from predators.
Female grouse nest and hatch their young during the
May- June period. Male grouse usually remain on their drumming
territories during this same time. Domey ( 1960 ) also found
that high temperatures in May resulted 5.n above average age
ratios and almost even sex ratios in fall. However, a cold
May resulted in low production and a loss in adult females.
High populations are less responsive to favourable May tempera-
tures than low populations.
(d) Summer brood co vers
Abandoned fields and clear areas adjacent to cover
provides a source of insect food. i^ienoptera and Coleoptera are
the more important groups of insects to grouse ( Edminster 1947 ).
Insects are primary contributors to the food supply of young
grouse from hatching time to mid- July, During July and August
a brushy understory is very useful to broods because it provides
summer shelter with abundant insect life close to the ground
( Bump 1947, Edminster 1954 ). Typical understory may consist
of shrubs and herbs such as striped maple, choke-cherry, red
osier, dogwood, jewel weed and various :ems, with sedges in the
more open areas. This type of cover provides a lower maximum
temperature to the broods than the more open cover types. During
the moulting period (July-August) grouse broods are reluctant to
flush ( Edminster 1947 ) . They tend to move into dense cover to
avoid flushing. Dusting sites are usually established on ex-
posed dry mineral soil or on dry rotted wood near old logs and
sturros. These areas are usually situated in such a position as
to take full advantage of solar radiation. Larsen ( 1958 )
found that warm days in spring and summer tend to be associated
with a high grouse population the following April and warm days
during winter tend to be associated with a low grouse population
the following April.
(e) Fall feed5_ng covers
Dennis ( 1966 ) found that during the September -
October period, due to the general availability of food and less
c::treme climatic conditions, grouse tend to distribute themselves
fairly uniformly in ail types of cover. During this period, the
diet usually consists of fruits and leaves ( Edminster 1947 ).
Birds tend to disperse as the brood structure breaks down in the
fall. During November and December there tends to be a general
regrouping of grouse activity occur ing from late fall to February.
The groups disband when spring territories are established.
Although the literature on Ruffed Grouse ecology
covers many areas of investigation, the aspect of microclimate
seems to have been barely scratched. Needless to say more exact
documentation on the microclimate of preferred cover types is
badly needed by game managers if this species is to be prooerly
Bump, G., R.W. Darrow, F.C. Edminster, and W.F. Crissey. 1947.
The ruffed grouse: Life history, propagation, management.
New York State Conserv. Dept. 915 nr> .
Dennis, D.G. 1966. Selection of cover types by ruffed grouse,
in Wellington County, Ontario. M.A. thesis, University of
Guelph. 84 on.
Dorney, R.S. and G. Kubat, 1960. Relation of weather, narasitic
disease and hunting to Wisconsin ruffed grouse populations.
Technical Bull. Ho. 20. Wise. Conserv. 63 pr> .
Edminster, F.C. 1947. The ruffed grouse , Macmillan Co.,
New York. 385 pp.
Edminster, F.C. 1954. American game birds. C. Scribner's Sons,
New York. 195 - 242
Geiger, R. 1965. The climate near the ground. Harvard Univ.
Press, Cambridge, Ilass. Revised Ed., 611 pp.
Grange, W.B. 1949. The way to game abundance. C. Scribner's
Sons, New York. 365 pp.
Gullion, G.W. 1961. A technique for winter trapping of
ruffed grouse. J. I/ildlife Management 25: 428-430.
Hungerford, K.E. 1951. Ruffed grouse oopulations and cover use
in northern Idaho. Trans. N.A. T :".ldl. Conf. 16:216-224.
Kittredge, J. 194G. Forest influences. McGraw-Hill Book Co.,
New York. 394 r>v>.
Larsen, J. A. and J.F. Lahey. 1958. Influence of weather upon
a ruffed grouse oooulation. Journal of Wildlife Mgt.
22(1): 63 - 70.
Marshall, W.H. 1965. Ruffed grouse behavior. Bio Science 15(2):
Palmer, W.L. 1961. A study of ruffed grouse drumming sites in
northern Michigan. Mich. Dept. Conserv., Game Div. Dent.
No. 2337. 40 pd.
Sparkes, C.H. and M.F. Buell. 1955. M5.croclimatological
features of an old field, and an oak-hickory forest
in New Jersey. Ecol. 36:363-374.
Spurr, S.H. 1957. Local climate in the Harvard Forest. Ecol.
A SURVEY OF WATERFOWL AND WATERFOWL HUNTING
ON WALPOLE ISLAND INDIAN RESERVE
J.M. COLLINS, Biologist
Ontario Waterfowl Research foundation
The major objective of this study was to obtain
the background information necessary to produce a management
plan for hunting on the Walpole Island Public Marsh. Waloole
Island Indian Reserve has an area of some 150 square miles
(Harrington 1965), which can be divided into three distinct
areas: the wooded area, primarily at the north end of the
island; the cultivated farmland, in the central portion of
the island; and some 20,000 acres of marshland, at the south
end of the Island.
Approximately 15,000 acres of the wetlands are
leased to six private hunting clubs. The remaining 5,000 acres
(the 'public Marsh 1 ) is not leased, but is managed by the
^Jalpole Island Indian Band. Public waterfowl shooting is pro-
vided there for non-Indians, who pay a fee for the privilege of
hunting, and who must be accompanied by an Indian guide while
hunting. The Public Marsh is centrally located in the midst of
the marshland, adjacent to the six large leased marshes.
A major drainage programme has been completed in
the last 10 to 15 years, converting about 3,000 acres of marsh
into farmland. Presently, the farmland is used almost exclusivel}'
for growing corn* A few acres of so3/beans are grown annually.
The 20,000 acres of marsh which remain undrained lie adjacent
and to the south of the reclaimed farmland. Large flocks of
ducks feed in the cc::nfields once harvesting is underway. This
is accepted as the. normal practice by the IJalpole residents,
indicating that the pattern has been established for some time.
Several persons did comment that the number of birds feeding in
the c ields has increased with the increase in the number of acres
o" corn being grown adjacent to the marshes. Birds in the r>ast
may have fed some distance inland, off the Island, but now can
concentrate on the newly farmed fields on the Island.
Work was begun in the r*ast T ew years to convert
the Public Marsh to a controlled water-level marsh. Dyking has
been done by drag- line and the fill has been levelled and
racked to create an access road to the edge of Johnston's Bay.
As r>art of the dyking operation, a ditch three to five feet
deep was dug around the Inside perimeter of the marsh, thus
providing a channel for boat travel. A ditch running east-
west was also dug b}' drag-line, to prov5.de access by boat to
shooting ponds located in the middle of the marsh.
The dyking operation is now complete exceot for a
section of approximately one and one half miles along the east
edge of Goose Lake, tihen this section is completed, it will be
possible to control the water level on approximately 2,500 acres.
Two small pumps, tractox 1 - driven, pump water from
the Johnston Channel into the marsh. Also, a larger pumo, driven
by automobile engine, has been installed next to the marsh, to
punm> water from the farmland into the Johnston Channel.
2 * HABITAT
The Public Marsh is at present densely overgrown
with emergent vegetation. There are relatively few areas of
o pen water. .As a result of the low water levels on Lake St.
Glair in the early 1960's, large portions of the marsh bottom
were exposed, and only shallow water oersisted on the remaining
areas. These conditions favoured the growth of cattails and
sedges and resulted in the abundant growth of vegetation that
now persists over most of the marsh.
The level of the lake has -risen, so that much of
the marsh is now covered with 18 to 24 Inches of water, with three
to three and a half feet of water in the ponds. There are two
<xcy areas in the marsh — one in the northeast and one in the south-
east- -that support mixed stands of grasses, sedges, goldenrod,
daisies, and other 'dry- land 1 slants. The wet portion of the
marsh is covered with stands o; T cattail, cattail-bulrush mixed,
bulrush, with some ohragmites ('quills 8 ) along the dyke edge.
3 • HATERFCftfL POPUL ATIONS
(a) Feeding p atterns
The most conspicuous pattern, of waterfowl behaviour
observed was the feeding activities of large flocks of dabbling
ducks, mainly mallards and blacks, with some pintails, in the
cornfields north and east of the Public Marsh. Throughout October
end November, large flocks (often totalling in excess of 10,000
birds) moved to the cornfields from the marshes twice a day --
in the early morning and late afternoon. Rain, wind or fog dis-
rupted this pattern somewhat. Then the birds moved to and from
the fields throughout the day.
The normal weather, flights from the marshes to the
cornfields began just before daxm and continued for about an hour.
By the end of the hour, birds started leaving the cornfields and
returning to the marshes. The large flocks left the cornfields
about two hours after sunrise. In the afternoon, there was no
'beginning time' for arrival in the fields. Rather, the number
of small flocks moving to the fields increased steadily through**
out the afternoon. Departure for the marshes after feeding was
sporadic until just before sunset. Then numerous flocks of
several hundred birds each left the cornfields together, breaking
up into smaller flocks as they approached the marshes.
A crew from the Canadian Uildlife Service (usually
including the author) conducted weekly aerial surveys of the
waterfowl populations on the Lake St. Clair marshes. Counts of
waterfowl were made at mid-day every Sunday throughout the hunting
season. During these mid-day surveys, few birds were seen on
the Public Harsh. Large concentrations of dabbling ducks (mostly
mallards and blacks) were however, present in the sanctuaries of
tltc neighbouring marshes, particularly St. .Anne's Marsh and the
marsh leased by Bradley Brothers.
The majority of the birds seen on these surveys were
in the resting or refuge areas, rather than in the feeding sanctu-
aries. Several marsh managers reported that little of the gram
put out in the feeding areas was being eaten, indicating that the
birds were obtaining sufficient food in the fields to satisfy
their nutritional requirements.
The hunting pressure and the progress of the corn
harvest had a direct e :fect in determining which fields were
utilized by the ducks. The first fields harvested (by picker and
sheller) were directly north of the Public Marsh. The
birds moved into these fields almost immediately, with the hunters
(Walpole Island residents) not far behind. The abundance of
hunters in the few fields yet harvested produced intense hunting
pressure with the result that most of the ducks were frightened
off. However, the com harvest proceeded at a rapid rate, each
day making more fields available for feeding. The ducks and
hunters, were then dispersed over a wider area, hunting pressure
decreased, and the number of b5.rds field feeding increased. This
in turn attracted more hunters into the fields .
At this time, the corn harvest began on St. Anne's
Island, north of St. Arme's Marsh. Because St, Anne's Island is
an island, it is inaccessible to most of the hunters. Consequently
the ducks quickly shifted to St. Anne's, where they could feed
with only a little disturbance. By the end of October, thousands
of ducks were feeding each morning in those fields. Heavy feeding
continued there until most of the dabblers migrated out of the
The shift in feeding sites, from Walpole to St,
Anne's Island, and the increased visibility given the ducks by
removal of most of the corn, drastically decreased the hunting
success in the fields. Many hunters lost interest. By November,
it was possible once again for ducks to feed relatively un-
disturbed in many of the fields north of the Public Marsh.
(k ) Flight patterns
The ducks which fed in the. cornfields were mallards,
blacks, and some pintails. These are the ducks preferred by the
hunters, i.e. the paying customers, so the 'lock movements of
these ducks is extremely important to hunting success. For ex-
ample, mallards and blacks made up 74% of the kill of all ducks
by non-Indian hunters, in 1968.
In the morning the ducks came to the fields from
every direction but the north. After feeding, they returned
to several distinct areas within the marshes, following one o :
three flight lines. In the afternoon, the same three flight
lines were followed, in the reverse direction, back to the
cornfields. However the flocks adhered to the flight lines less
than in the morning, small flocks 'wandering about' on the wa]
to the fields. After feeding in the afternoon, the birds
dispersed in many directions, toward the marshes, generally
ignoring the flight lines. Flight lines often shifted as a
result of wind, rain o:' snow, fog or hunting pressure in the
rarshes or the cornfields.
The important thing to note concerning these
flight lines is where they ended. At mid-day, the ducks sought
out a quiet undisturbed loafing area. Good loafing areas appear-
ed to be scarce, since most of the ducks could be found concen-
trated in a few acres. Xiring the two periods of field- feeding,
ducks were not so critical. They often returned to the same
cornfield for several successive days in snite of moderate hunting
pressure. This was particularly evident on one portion of St.
Anne's Island, where the ducks became especially unwary, probably
because of the extremely large flocks feeding there. In the
evening, the ducks were even less choosy about finding a resting
spot. Apparently 'any -?ort in a storm 1 applies as darkness
(c) Kinds and numbers of waterfowl
Almost every species of waterfowl found in eastern
North .America can be found in the Lake St. Clair area at sometime
in the fall. The fall of 1968 was no exception. The species most
commonly observed were mallard, black., widgeon, canvasback and
redhead. Because diving (or bay) ducks were virtually ignored by
most hunters, emphasis throughout this report is on dabbling
The weekly aerial surveys :;ave a grand total of
142,500 dabbling ducks counted on four Ualpole marshes, between
October 1 and December 16. The four marshes were St. Anne's
1-iarsh, the Smith or Bradley Marsh, the Anderson Harsh, and the
kr.blic Marsh. The counts for each marsh are given in Table I.
4 « T .MERF0WL HUNTI NG
Between October 11 and November 8, 1968, it was deter-
mined, by personally interviewing hunters and/or guides, the results
of 122 daily hunts. In 122 hunts, 500 ducks were killed, and 121
To determine the total number of ducks killed in the
season, the following calculations were made:
i. The total sale of daily hunting permits
was 150 o -ermits
ii. Each daily hunt bagged 4.1 ducks
iii. 1500 daily hunts 35 4.1 ducks - 6,150
iv. Each hunter with a season Dermit hunts at
least 10 times in the season. ( At $150
per season permit versus $15 per daily
permit, a purchaser of the season permit
plans to hunt at least 10 times. Some
likely hunt more than 10 times, a few
likely hunt less. Ten is a conservative
v. The holder of a season permit shoots the
same number of ducks per day (4.1) as the
holder of a daily permit
vi. There were 43 season permits sold in 1968
vii. 43 permits x 10 hunts per season x 4.1
ducks per hunt = 1763 ducks
viii. TOTAL DUCKS BAGGED = 6150 -f- 1763 = ... 7913 ducks
TOTAL DUCKS LOST = 1500 + 430 = ... 1930 ducks
ix. .Total number of hunts - 1930
The total of 9843 ducks killed is to be considered
a conservative estimate of the total kill for these reasons:
i. Most hunters were interviewed in the company of their
hunting companions, and/or guide, often as they filled out their
declaration slips permitting them to export their ducks through
customs. It is unlikely that any hunters claimed to have shot more
ducks than they actually did.
ii. A few hunters admitted to shooting over the limit of five
ducks, and giving the extra birds away. It is not known how many
others did likewise, and did not admit it. The number of addition-
al ducks killed this way is probably low. Several guides in-
dicated that they refuse to let their hunters shoot additional
birds because ducks #6 and #7 could be ducks #1 and i-2 for
iii. The crinkling loss (birds shot but not retrieved) is
probably in excess of the one duck per hunter day reported.
Hunters tend to report .ewer ducks lost than in fact were lost be-
cause of 'the wastefulness' asnect of losing ducks. Many of the
hunters interviewed 'couldn't remember' how many ducks the}'' had
lost. Several gave estimates lower than their guide later re-
Therefore, it nay be concluded that the total kill by
non- Indian hunters, on the Public Marsh and adjacent cornfields
I2E A Ji COUI-TT OF DA BBLING PUCKS O BSERVE D ON
FOUR SSIfflTED MARSHES (M IJALPOLE ISLAND ,
BETWEEN OCTOBER. 1 AND DECKMDER 16, 196"
NO. OF DUCKS
% OF TOTAL
DUCKS BAGGED PER HUNTER DAY CN FOUR PUBLIC
SHOOTING GROUNDS IN ONTARIO **
M- V ■ ■ ■III- ■ H ■ ■ ■ ~ ,., ,
** from annual reports, Ontario Department of Lands and Forests
was In excess of 10,000 ducks.
How does the hunting success here compare with
hunting success elsewhere? A success rate of 4.1 ducks per
hunter day when the bag limit is 5 is phenomenal. It compares
favourably with the success on the neighbouring marshes (approx-
imately 4.5), and contrasts drastically with success on public
shooting grounds elsewhere in Southern Ontario (Table II).
(k ) Reven ue cro m hunt ing
*• Ferrc it fees
Any non- Indian hunting on the Public Marsh
must have a Walpole Island Hunting Permit. There are two types
of permits; a season's permit which costs Si50, and a daily
permit which costs $13. The permit system was introduced about
1952, with a daily permit only, at a cost of $5 per day. A short
time later, the season permit, costing $100 was added and the
daily fee increased to $10. In 1967, the fees were again increased
to the present rates. The increase in permit sales, and revenue,
over the past 10 years is illustrated in Table III.
ii • Guide fees
Approximately 15 band members throughout the
hunting season devote their full time to guiding hunters in the
Public Marsh. In addition, there are 15 to 30 part-time guides
who are called upon when a regular guide has more hunters coming
on one day than he can handle.
The number of full- tine guides has remained
relatively constant for the past few 3'cars. A gentleman's
agreement has been effective in determining which areas of the
marsh are used by which guides. Most of the marsh has been
divided under this arrangement and any areas remaining unoccupied
are considerably inferior with regard to hunting success.
For a person entering the guiding business, a
fairly substantial initial investment is required for such items
as: boat(s), motor (s), decoys, waders, and raingear. Also, tools
and materials must be obtained for building blinds and cutting
shooting ponds. Time and labour must be spent in constructing
blinds, and contacts must be made with potential clients. These
factors plus the absence of more good hunting areas have served
to keep the numbers o r: guides relatively constant.
SALE OF WALPOLE ISLAND HUNTING PERMITS
•'-!- total estimated; 1,434 daily permits sold by end November
'} from Harrington, 1965
The daily fee for guiding services is set by each
individual guide, and varies somewhat among the various guides.
Many of the hunters interviewed had paid $20 per man per day.
Usually one guide accompanied two hunters. Some of the hunters
considered the guide fee they paid to be a personal matter, and
declined to give this information. Others indicated that they
had hired a guide on a seasonal basis, and could not ( or would
not) estimate the daily rate. None of the hunters Interviewed
thought the guide fee to be excessive.
Lacking more precise information than the above, $20
per man oer day is used as the average guide fee paid in 1968.
The income to all guides, before operating expenses are deducted,
is estimated as follows:
1500 daily hunts at $20 ner hunt $30,000
430 hunts on season permits at $20 . . 8,600
TOTAL GROSS REVENUE $30,600
iii. Total revenue from hunting, 1968
Permit sales $20 , 950
Guide fees 30,600
( c ) Efficiency of the present hunt inn; methods
There appears to be three possible methods
of conducting the hunt on Jalpole Island:
1. Attract a large number of hunters, each paying
a large sum of money for a day's hunt.
2. Attract a smaller number of hunters, each
paying a large sum of money for a day's hunt.
3. Attract a large number of hunters, each paying
a small sum of money for a day's hunt.
Method 01 i;?ould obviously bring the greatest financial
return to the Band. However, a duck hunter must receive a duck
hunt worth a lot of dollars to induce him to pay a lot of dollars.
Ouch a hunt is carried out in an undisturbed manner, without inter-
ference from other hunters, and with an abundance of ducks, and
ample shooting opportunities. But duel: hunting is such that it is
impossible to have this type of hunting when large numbers of
hunters are using an area. Therefore, method #1 is not possible.
Method (rl is presently being used, and it is
successfully providing a considerable anount of revenue to the
Walpole Indian Band, Its continuation denends on the Public
Harsh continuing to provide a hunt worth a lot of dollars. If
the quality of the hunt declines, so will the number of hunters.
Other hunters, who are less interested in the quality of the
hunt, will be required to replace the lost revenue. These hunters
are rarely prepared to -oay much money Tor the opportunity to hunt,
so that large numbers of hunters are necessary to produce the same
revenue. Method #3 would then be operating. However, it is
nrobably impossible to attract the larcge numbers of hunters re-
quired for method #3 to be able to compete financially with method
For example, in 1963, some 2,000 hunters paid
$28,950 for permits to hunt on Walpole Island, and $38,600 for guide
services. This amounts to approximately $35 per man for one day's
hunt. At $10 per man nev day, almost 7,000 customers would have
been required to provide the same revenue. In a duck season ten
weeks long, this would mean 700 hunts per week. At $5 per man r>e?:
day, 1400 hunts per week would have been required.
The decrease in the hunting success with 700 hunts
per week would be so drastic that it would be most difficult, if
not impossible, to induce 7000 hunters to pay $10 for a day's
hunt. It would be especj.ally difficult when waterfowl hunting is
available elsewhere, on provincial!}' operated units, for $4 per
blind per day, two men to a blind. Therefore it may be concluded
that the present method, whereby a moderate number of hunters each
pay a considerable sum of money to hunt, is the most efficient and
profitable means of providing revenue to the Band.
( d ) Factors affect ing present h untin g operations
Since method #2 is the best method, it is important
to understand the factors that allow it to operate so successfully.
The primary factors are:
i. Hunting success
ii. Number of ducks in immediate area
iii. Cost of hunting in the Public Marsh
iv. Frosrfjnity of Walpole Island to the Detroit-
Grosse Pt. - St. Clair Shores complex
v. Difference in bag limits between Ontario and
i. Hunting success;
The outstanding feature of hunting on Walpole Island is the
success each hunter lias, day in, day out, week in, week out.
This is what the hunters rave about ...... 'best place in North
America, can come anytime and shoot a limit'.
ii . Number of du cks in the im mediate area:
There are tremendous numbers of ducks in the Lake St. Clair
area in the fall, so that the hunters can be selective as to the
kinds of ducks they shoot. Of the ducks shot on the Public Marsh,
58% were mallards, whereas mallards composed only 28% of the total
waterfowl kill in Ontario in 1968. The continuing high rate of
hunting success throughout the fall is clue, in a large part, to
the continued presence of large numbers of ducks until late in
i ii. Cost of hunting in the Public Marsh
The high cost of a day's hunt contributes directly to the success
of the present system, for several reasons. The cost prohibits
all but a few individuals from hunting on Walpole Island. Because
only a few persons can afford to hunt, the hunting pressure is
moderate, and evenly distributed throughout the season. With only
a few people hunting, the pressure on any given day is low. Host
of the hunters, however, return several times in the season, so the
total number of hunts is moderately high and a good revenue is
Distributing the hunting pressure over the entire season helns
to maintain the high level of hunting success. This method
contrasts with the 'opening day' syndrone that occurs throughout
the rest of the province when approMirmtoly 75% of the season's
kill is taken on the first two weekends, and local duck populations
are often 'burned out*.
The cost of hunting on Walpole Island includes payment for the
services of a guide. The present guide fees, while adding consider-
ably to the cost of hunting, are attracting excellent guides. The
presence of a guide in the blind contributes in a large way to the
hunter's success, and his subsequent return at a later date. The
presence of a guide accomplishes what a long list of rules and
regulations attempts to do on government controlled waterfowl
units, and does it in a friendly way. The guides are in many ways
responsible for the quality of hunting in the marsh.
iv. Proximity of Walpole _Island to the Det roit area :
With cost playing such an important role in the system, it
is obvious that the operation is successful as a business venture
because it has a wealthy Dooulation from which to obtain its
customers. The 1968 permit sales showed that 7% of the hunters
were from Ontario, but 93% of the hunters were American; 74% were
from Michigan, 12% from Ohio, and 7% from the remaining states.
Almost all of the Michigan hunters came from the Detroit-Grosse
Point - St. Clair Shores area, an hour J s easy drive to Walpole
Island. The appearance, attitudes, occupation, and home address
identified the hunter as having a substantial income.
v. Difference in bag l imits be tween Ontario an d Mic higan:
Many hunters expressed dissatisfaction with the short season,
and the low daily limits in effect in Ilichigan. For example, the
196G season ran from October 10 to November 8. The daily bag limit
was 3 ducks, not more than 1 mallard, 1 black duck, 2 wood ducks,
1 canvasback or 1 redhead. The increase in the sale of Walpole
Island permits is undoubtedly due to the longer season and more
liberal daily limits in Ontario. The 1968 season opened in south-
em Ontario on October l j and closed on December 14. The daily
bag limit was 5 ducks, not to include more than 2 canvasbacks or
2 redheads, or more than 4 wood ducks.
The bag limits in Michigan are not likely to be increased to
five in the foreseeable future. Ontario limits may go down, but
will likely remain above the Michigan Limits. As long as there are
duck hunters in Michigan, they will tend to be drawn to Walpole
Island. However, it is possible that the bag limits can become
reduced to the point that hunters will give up hunting, particularly
if costs are considered to be unreasonable.
In summary, the present operation is successful as a business
venture because it has a wealthy population from which to obtain
its hunters, and the hunters that come to Ualpole Island go home
satisfied. The moderate hunting pressure, evenly distributed
throughout the season, an abundance of ducks, and the high quality
of hunting resulting largely from the guides' supervision, Droduce
a satisfied hunter.
(e) Atti tudes of the hunt ers
In any other part of Ontario, the lack of customers
able to r>ay the high cost of hunting would likely cause this tyr>e
o :■:" hunting business to fail. It can fail here as well, if the
customers become dissatisfied and consider the costs unreasonable.
Steps must be taken therefore, to ensure that hunters are
One source of dissatisfaction that now exists, is
the cost of the Walpole Island permit. Many of the hunters in-
terviewed considered the $15 fee to be excessive. The degree of
dissatisfaction varied, however, with the hunting success that the
hunter had on the clay he was interviewed. Fifteen dollars for
five ducks is easier to accept that fifteen dollars for one duck.
The reluctance to pay the Walpole Island fee
might be overcome by indicating to the hunter what he is receiving
for his money. The same person who complained that the permit
fee was excessive usually was satisfied with the guide fee, which
generall} 7, exceeded fifteen dollars. However, the hunter can see
what he is getting from his guide a boat, motor, gasoline,
decoys, a blind, a 'private 1 pond, and ducks . The return to the
hunter from his permit fee is much less obvious. In fact, many
hunters questioned whether they received anything in return for
the money spent.
I consider that the permit fee is each hunger's
'rent payment 1 to retain the marsh as a hunting marsh. In this
sense, the Band in 1968 received approximately $30,000 in rental
money, in exchange for the use of the marsh by the public for duck
hunting. I believe that the rent concept would explain to the
hunters why there is a fee, and that it is worth making this idea
known to the hunters.
A potential source of dissatisfaction, and one that
has apparently caused problems in the past, is having too many
hunters in the marsh at one time. Too many customers is a problem
a lot of businesses would like to have, but in this case, too many
customers can destroy the product being cold, and consequently
produce a lower revenue. Walpole Island is selling a specialised
product -- top quality hunting — to a small group of people --
those prepared to hunt several times in a season, at $35 a day.
The present financial success o.c the hunting busi-
ness is a result of attracting 'repeat customers'. In the 196G
season, hunters holding daily permits hunted an average of 2.5
times on Walpole Island. Most hunters have been coming to Walpole
for several successive years. The new customers had usually learn-
ed of hunting on Walpole Island by word- of -mouth, or by accompany-
ing a friend who had hunted on the Island previously. Thus, losing
one customer can significantly affect the level of revenue,
because his repeat business is lost, as well as that of his hunting
During the early 1960 's, which were years of
low water levels on Lake St. Clair, large portions of the marsh
were dry and unsuitable for duck hunting. The hunters were
forced to hunt in a greatly reduced area. The decrease in permit
sales in 1964 was probably a result of hunters being dissatisfied
with the quality of the hunting under crowded conditions.
5 * ROLE OF THE PRI VATE M ARSHES
If the private marshes of the Lake St. Clair region
ceased to exist, the large concentrations of waterfowl there would
become a thing of the past. The private marshes are probabl}/ best
described as 'refuges on which some hunting takes place 1 . Personal
interview with marsh managers, and a report by Bryant (1965) in-
dicate that hunting pressure on the marshes is light. It is
estimated that in 1968 the total bag for all private marshes in
the Lake St. Clair area was approximately 10,000 birds, compared
with an estimated bag of 3,000 birds on the Public Marsh. Aerial
surveys revealed that the private marshes are used as rest:.n~ areas
by large numbers of migrating waterfowl (Table I). All of the
private marshes on Ualpole Island have feeding sanctuaries, and
most have undisturbed refuge areas as well, which attract and hold
the birds. Thus these large, lightly hunted marshes act as reser-
voirs and provide birds for hunting on the adjacent Public Marsh.
The sporting tradition associated with these
private clubs, most of which have a long history, has given the
members a strong conservation conscience. The rules regulating
the hunt are often severe. St. Anne's marsh probably typifies
the leased marshes on Ualpole Island in this respect. St. Anne's
lias two feeding sanctuaries, and two refuges. The maximum number
of hunters on one day in this 7000 acre marsh is eight, x*ith five
the average. The marsh is hunted only four days a week; Tuesday
afternoon, Wednesday morning, Friday afternoon, and Saturday
morning. To reduce the number of crippled ducks, hunters do not
shoot at flocks of ducks larger than five. The total bag runs
about 1500 ducks oer year. Only 1500 ducks were shot, but 73,500
dabblers and 108,000 divers were counted using the marsh this
past fall. The hunting clubs have in effect created waterfowl
refuges, and what is amazing is that they are willing to pay a
considerable amount of money for the privilege of doing so.
It cost the average hunter of the private marshes on
Lake St. Clair $57 to bag a duck in 1964 (Bryant 1965). Obviously
the hunter is buying more than a duck. The amount a hunter will
spend reflects not the value of the marsh, but the value of the
hunting on that marsh. To attempt an economic comparison of
marshland to corn land, and ducks to corncobs, is to misunder-
stand hunting values.
At the time of lease renewal, it is most important
to consider the Conservation history 1 of the club(s) concerned.
How much has the club contributed to the value of the hunting on
that marsh? To the best of the author's knowledge, the clubs have
all been good tenants, in spite of the fact that most of the leases
do not include •performance 1 clauses.
The suggestion has been made that performance
clauses should be included in any new leases, to protect the water-
fowl population from over-hunting, by ensuring the continued good
behaviour of the lessees. Such clauses would include an upper
limit to the number of hunter-days allowed on each marsh in a
season, might possibly set a limit on the number of ducks killed
annually on each marsh, and possibly set down guidelines for
management of the marshes. In my opinion, such clauses could not
accomplish their intended purpose, and could have a detrimental
effect on the waterfowl if the goodwill of the club members is
lost. It is one thing for the clubs to voluntarily maintain the
marshes as 'refuges 1 , but quite another thing to require them to
do so. It would be unrealistic to restrict the clubs to the
present total kill, considering the money the clubs spend on
management each year. For the money spent, they are entitled to
more ducks than they arc presently shooting. Therefore, to be
acceptable to the clubs, the limits would have to be above the
present total kill. If the clubs were inclined to shoot to the
limits, the kill would increase beyond the present level.
Second, a restriction on the total kill, or the
hunting pressure (in hunter-days) would be almost impossible to
enforce. Who is going to count the ducks shot and the number o^
hunters hunting? If the clubs are ejected to police themselves,
why not trust them to manage the resource as they have been
doing? It is wishful thinking to believe that merely passing a
by-law will make people law-abiding, if they haven't been in the
Third, any restriction on total kill and hunting
pressure that is liberal this year might well be harmful five
years from now. Waterfowl populations can fluctuate drastically
from one year to the next. For this reason, federal waterfowl
regulations concerning hunting are set annually, to adjust ~or
these fluctuations in numbers of waterfowl. It would be most
difficult to insert meaningful limits in a long term lease.
Fourth, the proposal to set down guide-lines for
marsh management assumes a greater knowledge of management tech-
niques than actually exists. In fact, many of the local club
managers knoitf more about marsh management than many waterfowl
'experts 1 . The managers are open-minded with regard to suggestions
for improving their marshes, provided the suggested techniques
work. Guide-lines are not really necessary. Unfortunately, tech-
niques with predictable results have not been developed as yet.
The final argument against performance clauses is
the performance of the clubs in the past, without any of these
clauses. It has not been by accident that the clubs have created
'refuges with some hunting'. The quality of hunting demanded by
a person paying $57.00 to bag a duck regulates the hunting be-
haviour more effectively than by-law ever could. There is of
course one important, although unwritten, performance clause in
every marsh lease, even now. When a lease expires, the Band
Council can refuse to renew the lease if the performance of the
lessee in the past has not been satisfactory. This is a two-way
street, however. If unreasonable demands are made of the lessee,
he may refuse to renew the lease. Such action could make it
difficult to attract new tenants. Any prospective tenants are
certain to determine the reason for the lease not being renewed,
and they are likely to consider unreasonable demands made in the
past as an indication of the future.
6 . recohiendatioe :
The present public hunting operation on VJalpole
Island is efficient and profitable for the Band. Major changes in
the system are not necessary. However, several steps should be
taken to ensure that the system continues to be successful. Too
many hunters in the marsh is likely to become an immediate problem.
The number of hunters is now approaching the point where the
quality of the hunting may begin to decline.
Additional hunters could be accomodated successfully
if more of the present marsh were utilized. Plight now, the north-
east and south-east sections of the marsh are of little use to ducks
(and therefore duck hunters), as those sections are high and dry most
of the season. Flooding of the area, by pumping is not feasible.
Putting enough water on this area to make it useful, would put too
much water on the rest of the marsh. Enclosing the area with its
own dyke is possible but expensive. The additional cost would not
likely be matched by an equal increase in revenue.
The best solution for making this area useful is to
create a series of shallow ponds either with a drag- line, or by
blasting with ammonium nitrate. Extra hunting would result, but
in addition, pot-holes in a grassy area such as this attract
nesting ducks, and could help increase production.
For these ponds to be used for hunting, it is im-
portant that the depth be controlled. fonds six feet deer- cause
problems when you are wading, and you are only 5' 6" tall. Maxi-
mum depth should be approximately 3 veet .
A second improvement that should be made in the marsh,
is to establish water-level control, by finishing the dyking ^re-
ject. This entails running a dyke from Dynamite Cut north along
Goose Lake and then connecting to the existing dyke. To minimize
costs of moving equipment, the drag-line might be used while it is
still at the marsh to dig shooting ponds, in the dry areas.
The new dyke and ditch should be built in such a way
as to prevent their being used as a new access route to the marsh.
Much of the hunting depends on the flight patch which brings the
ducks over the middle of the marsh. Travel back and forth along
Goose Lake could disrupt this pattern. Going one step further the
road should be closed on the dyke that runs east-west to the Romal
pump. This would remove most of the traffic and the resulting
disturbance along the north edge of the marsh. Consideration might
also be given to establishing a 'no hunting zone' in the cornfields
directly north of the marsh, where the ducks could field feed un-
disturbed. Farming would proceed as normal but no hunting would be
allowed. Two purposes could be served by such a refuge. First, it
would provide a 'safe 1 area on WalpoXe Island, so that all the birds
could not shift to St. Anne's Island to field feed. The refuge
would keep birds closer to the Public Marsh, and create another
flight over the marsh as the birds leave the fields, and move to
the loafing areas. The second purpose would be to move the. east-
west flight line closer to the marsh. Presently, hunting 5ji the
fields directly north of the marsh pushes the east-west flight line
north, away from the marsh. A 'buffer strip 1 between the marsh and
the zone of field shooting should overcome this problem.
The ability to control water-levels will overcome
the Droblem encountered in the Dast, of the marsh 'shrinking' during
years of low lake levels. But equally important, adjusting water-
levels in the marsh is one way of changing the kinds, and amounts,
of vegetation growing in the marsh. Therefore, plans for dyke con-
struction and pump installation should include the means for getting
water off the marsh as well as on.
Another suggestion concerning efficient use of
the marsh, is to create more shooting ponds over the entire marsh,
not just in the dry parts. More openings would mean that each
guide could have several ponds. By alternating shooting from
pond to pond, no one pond would be shot out.
While the steps just mentioned should permit more
hunters to hunt without affecting the quality of hunting, the basic
problem is merely postponed. It is almost certain that some day,
in the not- too-distant future, there will be too many hunters for
the public Marsh. The first, and most urgent sten to be taken to
cope with the problem, is to determine how many are too many.
This question must be answered soon, otherwise the answer may be
learned too late, when hunters do not return, and the revenue has
One very important clue to hunter satisfaction is the
number of times a hunter returns in each season, and the number of
hunters coming bach in successive years. When repeat business
starts to decline, something is wrong. Consideration should be
given to designing the records of the sale of hunting permits in
such a way that information about repeat customers can be readily
An attempt should also be made to develop a system
to measure hunter success, throughout the season as well as from
season to season. The declaration forms, which must be presented
to customs when ducks are exported through to the U.S. are one
potential source of this information. Another possibility is to
design a hunting permit with a tear-off portion on which the hunter
records his daily kill. The records would then be deposited at a
centrally located collecting station. The name of the guide should
also be included, since the guides could check the truthfulness of
the records being submitted.
The causes of a decline in the hunter success from the
present high rates should be examined very carefully. If there is
any indication that excess hunting pressure is responsible, correct-
ive steps should be taken immediately.
There appear to be several ways of coping with * too
many hunters 1 . One way is to expand -oubiic hunting into one of
the marshes presently leased to a private club. To lustily this
move, the increase in income from public hunting will have to at
least equal the money now received from the club including rental
payments and wages paid to guides. The ei^ansion of public hunting
into one of these lightly hunted nrivate marshes could drastically
upset the waterfowl movements that presently make hunting on the
lublic Marsh so successful.
A second possible solution to the crowding problem
is to make greater use of the field shooting opportunities. It is
difficult to predict precisely the areas the ducks will choose
each day for feeding. The resulting all-or-none aspect of field
shooting apoeals to a few hunters but most of the Walpole hunters
reject it for that very reason. They want their ducks. There-
fore, field shooting has only a limited potential as a solution to
A third, and probably the most unpopular solution is
to set a hunter-day quota for the Public Marsh. I expect that
excess hunting pressure will appear first on the few 'special 1
days in the season — opening day (for example, Oct. 5, 1963),
and Thanksgiving weekend, if the weather is right. Then most good
Saturdays will be busy. Finally, to escape the crowd on the week-
end, more hunters will come in mid-week. The quota system could
therefore be introduced in stages:
i.e. no more than X hunters in any one day
no more than Y hunters in any one week
no more than Z hunters in a season
With the number of prospective hunters that are likely
to come to Walpole Island in the future, this is to me, the only
The waterfowl resource, unfortunately, is not limit-
less. With the continuing decline in the number of mallards, it
would be short-sighted to expand the Public Hunting area, and
double the mallard kill on the Island. A quota on hunters will
help to protect the resource, and the quality of the hunting. A
resource which contributes so greatly to the revenue of the Band
deserves to be protected.
There is a resource which, as a potential source of
revenue, has not yet been utilized. With proper management, it
is possible that Walpole Island could support a large natural
population of pheasants. Evidence is that even now there is a
significant population of bob-white quail. Host c£ the duck
hunters interviewed were enthusiastic about the possibility of
hunting pheasants on Walpole Island. The customers are there.
Good quail hunting would be certain to attract those hunters who
own bird dogs, since there are few such hunting opportunities else-
where in Ontario, or I'ichigan.
It is imperative however , that the hunting be based
only on natural population. liaising and releasing birds would be
too expensive to be profitable. The quality of hunting released
birds is definitely inferior to hunting wild birds. I suggest
that like duck hunting, the key to success would be in providing
top quality hunting for the top dollar.
Pheasant hunting offers the advantage of a flexible
open season, x^hich can be adjusted to suit the local conditions.
It is possible that pheasant hunting could be used to attract
hunters to the Island for a different period of time than the 10-
week duck season. It night then become profitable for the Band
to consider settling up a T club-house-motel-restaurant 1 business,
vjfhen asked to suggest improvements in the hunting, almost every
hunter requested 'a -olace to meet other hunters', or *a place to
have a drink 1 , or 'a nlace to get a good dinner', or a 'good motel 1 .
These hunters are wealthy. They have money to spend. They should
be given the opportunity to snend more of it on Ualpole Island.
The financial success of the hunting operation is due
to the unique situation of the Public Marsh. It is located in a
major rest area for migrating waterfowl. It i;; surrounded by
'refuges' and is only a short distance from a large population of
wealthy sportsmen. An abundance of ducks is available. The
quality o; hunting is superb. Hunters realize this and are will-
ing to pay in proportion to the quality. Loss of high quality due
to crowding and excess shooting could have a deleterious effect
on financial success. Steps to cope with this problem should be
taken before it occurs.
Bryant, J.E. 1965. Great Lakes water levels and migratory
waterfowl, Ontario. A brief to the International Joint
Commission on Great Lakes Level. Hiraeo report, -ilea
with the Canadian Uildlife Service. 7 r>r>.
Harrington, C.H. 1965. An economic survey of the Walooie
Island Indian Reserve. Mimeo report. 67 pp.